A new key for the species of Ateuchus Weber (Coleoptera: Scarabaeidae: Scarabaeinae) occurring in Mexico, with a description of the first North American inquiline species from a rodent burrow (Rodentia: Geomydae) and new distribution records

RevistaBrasileiradeEntomologia62(2018)131–134

REVISTA

BRASILEIRA

DE

Entomologia

Systematics,

Morphology

and

Biogeography

A

new

key

for

the

species

of

Ateuchus

Weber

(Coleoptera:

Scarabaeidae:

Scarabaeinae)

occurring

in

Mexico,

with

a

description

of

the

first

North

American

inquiline

species

from

a

rodent

burrow

(Rodentia:

Geomydae)

and

new

distribution

records

Bert

Kohlmann

_,

_Fernando

_Z.

_Vaz-de-Mello

b_{UniversidadeFederaldeMatoGrosso,InstitutodeBiociências,DepartamentodeBiologiaeZoologia,Cuiabá,MT,Brazil}

a

r

t

i

c

l

e

i

n

f

o

Articlehistory:

Received10November2017 Accepted18January2018 Availableonline13February2018 AssociateEditor:AdrianaMarvaldi Keywords: Internalsac Neotropicalregion Scarabaeoidea Taxonomy Veracruz

a

b

s

t

r

a

c

t

ThefirstAteuchusWeber(Scarabaeinae)species,A.tuzasp.nov.,fromarodentburrowfromNorthAmerica isdescribed.Diagnosticcharactersarepresented;photographsofanadultmaleandillustrationsofmale genitaliaareincluded.AkeyforallknownMexicanAteuchusWeberspeciesisprovided.Ateuchushornai (Balthasar)isrevalidated.NewdistributionrecordsfromMexicoandCentralAmericaarereported.

©2018SociedadeBrasileiradeEntomologia.PublishedbyElsevierEditoraLtda.Thisisanopen accessarticleundertheCCBY-NC-NDlicense(http://creativecommons.org/licenses/by-nc-nd/4.0/).

Introduction

ThisworkisacontinuationofthestudyofthegenusAteuchus WeberforNorthAmerica,firststartedin1981(Kohlmann,1981, 1984; Kohlmann and Halffter, 1988), and thereafter extended to Mesoamerica (Kohlmann, 2000) and Costa Rica (Kohlmann, 1996–1997;KohlmannandSolís,2009,2012).Itdescribesthefirst knownAteuchus speciesfrom theburrowof a rodent collected inNorthAmerica.Ateuchushornai(Balthasar)isrevalidated.This descriptionandthepresentrevalidationincrease thenumberof previouslyknownspeciesfromMexicotofourteen.Asa compar-ison,twelvespecies areknownfromCostaRica(Kohlmann and Solís,2012).ThisstudyalsopresentsakeyfortheMexicanAteuchus species.ItalsoreportsnewAteuchusrecordsfromMexicoand Cen-tralAmerica.

Materialsandmethods

Mostspecimens studied came fromtheEntomology Section ofthe ZoologicalCollection of theUniversidadeFederaldeMato

∗ Correspondingauthor.

E-mail:bkohlman@earth.ac.cr(B.Kohlmann).

Grosso, Cuiabá, Brazil (CEMT) and the Entomological Collection at the Institute of Ecology (IEXA), Xalapa, Mexico. Other col-lections are cited only once and so no acronyms are used for them.

ThespecimenswerestudiedusingaZeissStemi2000-C stere-ozoom binocularmicroscope. Measurementswere madeto the nearest 0.1mmusing anocularmicrometer.Genitaldissections andpreparationsweredonefollowingthetechniquesdescribedby Zunino(1978).Genitalstructureswerestoredinmicrovialswith glycerin.Thestructuresoftheinternalsacweredrawnusingthe freehandtechnique.PhotographsweretakenwithaZeissAxioCam ICc1 camera.PhotographswereprocessedusingtheZeiss Axio-Vision 4 image processor. Morphological nomenclature follows Kohlmann(1984,2000)andGénier(2015).

Results

KeyfortheidentificationofAteuchusWeberfromMexico (modifiedfromKohlmann,1981,2000)

https://doi.org/10.1016/j.rbe.2018.01.002

0085-5626/©2018SociedadeBrasileiradeEntomologia.PublishedbyElsevierEditoraLtda.ThisisanopenaccessarticleundertheCCBY-NC-NDlicense(http:// creativecommons.org/licenses/by-nc-nd/4.0/).

132 B.Kohlmann,F.Z.Vaz-de-Mello/RevistaBrasileiradeEntomologia62(2018)131–134 1.Eyesviewedfromabovesixtimeslargerthanwide...2

–Eyesviewedfromabovetwoorthreetimeslargerthanwide... 4 2.Anteriorpronotaledgewithcompletemargin;baseoftheheadwitha wrinkledarea(Fig.3)inthecenter;sternellumformingacarinaalongthe middle;inquilineofrodentburrows.Mexico(Veracruz)...A.tuza KohlmannandVazdeMello,sp.nov.

–Anteriorpronotaledgewithincompletemargin;baseoftheheadwithno wrinkledarea;sternellumnotformingacarinaalongthemidline;

non-inquilinespecies... 3 3.Ovoidbody;pronotalpuncturesformingasemi-circleatthemiddleofthe pronotalbase;internalsacwithaverydevelopedspinyfasciaandthree copulatoryspicules;puncturesofthelowersurfaceoftheanteriorfemorafine anduniformlydistributed.MexicotoPanama(SinaloatoAzueroPeninsula)... rodriguezi(PreudhommedeBorre)

–Longbody;pronotalpuncturesnotformingasemi-circleatthemiddle partofitspronotalbase;internalsacwithaweaklydevelopedspinyfasciaand twocopulatoryspicules;puncturesofthelowersurfaceoftheanteriorfemora moderateinsizeanduniformlydistributed.Mexico(Higherreachesofthe BalsasRiverValley;Guerrero,Morelos,andPuebla) ... halffteriKohlmann 4.Apexoftheelytralstriaeweaklyimpressed...5 –Apexoftheelytralstriaestronglyimpressed...6 5.Shagreenelytra;internalsacwithaverywelldevelopedfascia;pygidium veryconvexandshagreeninitsupperpart.MexicotoColombia(Oaxacato Chocó)... candezeiHarold

–Elytranotshagreen;internalsacwithaweaklydevelopedfascia;pygidium flatandnotshagreen.MexicoandBelize(Atlanticslope;Chiapas,Campeche andOrangeWalk)...laetitiaeKohlmann 6.Lengthlargerthan8mm,andventralsurfaceofanteriorfemurveryfinely punctatethroughout,lackinganylargerorcoarserpunctures(onlyonefemale known,8.4mm,Necaxa,Puebla)... hornai(Balthasar) Eitherlengthshorterthan8mmorventralsurfaceofanteriorfemurwith areasoflargerorcoarserpunctures...7 7.Ventralsurfaceofanteriorfemurfinelypunctatethroughout,fewlarger puncturesalongposterioredgeinsomeTexanindividuals;spermatheca taperingfromtheaccessoryglandinsertiontoapex.USAandMexico(Texasto Veracruz)...texanusRobinson

–Ventralsurfaceofanteriorfemurcoarselypunctatethroughoutoratits posterioredgeorapex;spermathecauniformlythickthroughout.Mexicoto Honduras... 8 8.Ventralsurfaceofanteriorfemurandpygidiumwithcoarsepunctures throughout... 9

–Ventralsurfaceofanteriorfemurwithfinepuncturesthroughoutorcoarse puncturesatitsapexoratitsposterioredge;pygidiumfinelypunctate...10 9.Spermathecaring-likeatitsbase;primarycopulatoryspiculefine.Mexico (PacificslopeoftheTransverseVolcanicSystemandSierraMadredelSur; JaliscotoOaxaca)... carolinaeKohlmann

–Spermathecanotring-likeatitsbase;primarycopulatoryspiculemassive. Mexico(TransverseVolcanicSystem;MichoacántoVeracruz)...klugi(Harold) 10.Anteriorpronotalmarginalmosteffaced;pronotalsurfacemoderatelyto coarselypunctate;pygidiumveryconvex;oneofthecopulatoryhooksofthe internalsacisalongandslenderfilament;oneoftheapicallamellaeofthe internalsachasacanoe-likeform.MexicotoNicaragua(ChiapastoMasaya) ... guatemalensis(Bates)

–Anteriorpronotalmargincomplete;pronotalsurfacefinelypunctate, sometimeswithsomecoarsepuncturesliningthebaseandmarginsofthe pronotum;pygidiumflattoconvex;noneofthecopulatoryhooksofthe internalsacisfilament-like;noneoftheapicallamellaeoftheinternalsacare canoe-like.MexicotoHonduras...11 11.Smallspecies(4.8–5.5mm);secondarycopulatoryspiculesoftheinternal sacverysmall.Mexico(VeracruztoYucatán)...perezvelaiKohlmann

–Bigspecies(6–8.5mm);sizeofthesecondarycopulatoryspiculesofthe internalsacsmalltobig.MexicotoHonduras(TamaulipastoLaPaz)... 12 12.Copulatoryspiculesoftheinternalsacsubequalandspine-like;nospiny faciespresentaroundthecopulatoryspicules.Mexico(Chiapas)...gershensoni Kohlmann

–Secondarycopulatoryspiculesoftheinternalsacsmallerthantheprimary one;spinyfaciespresentornotaroundthecopulatoryspicules...13 13.Elongate,convexandbroadbody(4.0–4.5mm);eyesdorsallytwotimes longerthanwide.Mexico(TamaulipastoChiapas)...illaesum(Harold)

–Oblong,veryconvexandbroadbody(5.0–5.5mm);eyesseendorsally threetimeslongerthanwide.MexicotoHonduras(ChiapastoLaPaz) ...chrysopyge(Bates)

Figures1–3.Ateuchustuzasp.nov.;1,dorsalhabitusofholotype;2,ventralhabitus

ofholotype;3,wrinkledareaatthebaseoftheheadofamale.

AteuchustuzaKohlmannandVazdeMello,sp.nov.

(Figs.1–4)

Typelocality:MEXICO:VERACRUZ,DosAmates(Catemaco),elev.

300–600m(18◦2921N,95◦0335W)

Diagnosis.ThisspeciesisdistinguishedfromotherAteuchusspecies

bythefollowingcombinationofcharacters:Longandslenderbody; dorsum glossy;clypeus bidentate; headsimply convex,lacking carinaortubercle;baseoftheheadwithasmallwrinkledareainthe center;anteriorborderofpronotumcontinuous,posteriorborder ofpronotumwithanareaofdenseandcoarsepuncturesmedially; elytrallateralportionrounded;sternellumformingacarinainthe middle;pygidiumslightlyconvex,lightlyshagreen,and inconspic-uouslypunctate.

Description.Holotypemale(Figs.1–4).Body.Elongate(Figs.1–2),

convex dorsally(Fig. 1).Size. Total length 10.0mm. Maximum width5.5mm.Color.Darkreddishbrowntoblack,lacking metal-lic sheen. Head. Clypeal margin anteriorly broadly V-shaped (Figs. 1–2); anterior margin slightly upturned, gena and frons coarselywrinkled,vertexcoarselypunctate,baseofheadwitha

B.Kohlmann,F.Z.Vaz-de-Mello/RevistaBrasileiradeEntomologia62(2018)131–134 133

smallwrinkledareainthecenter(Fig.3),eyesviewedfromabove sixtimeslongerthanwide.Pronotum.Transverse,strongly con-vex,surfaceglossy;anteriorpronotalmargincomplete;midline weakly impressed at base; pronotal surface punctate through-out, a group of coarsepunctures at thecenter of thepronotal base(Fig.1);lateralpronotalfossaeshallow.Elytra.(Fig.1).Striae fine, shallowly impressed on disk, becoming well defined and deeplyimpressed onapicaldeclivity;strialpunctures elongate-crenulate,slightlywider thanstria, separatedby2–3diameters ondiskandapicaldeclivity. Interstriaeslightly convexondisk, surfaceminutelypunctatethroughout.Thoracicsterna.(Fig.2). Proepisternumexcavateanteriorly,surfaceofexcavatedportion granulate,withfine andrather longsetae,borderedposteriorly by a well-defined keel. Proepimeron finely wrinkled. Sternel-lumglabrous andformingakeelatitscenter.Mesometasternal suturearchedmedially,marginalbeadfine.Mesosternumcoarsely puncturedformingrugulae.Mesepisternumflat,surface-forming rugulae. Metasternum evenly convex, disk evidently punctate, surfaceglossybetweenpunctures,laterallobesformingrugulae; midlineclearlyimpressedalongthreefourthsofitslength.Legs.

(Figs.1–2).Foretibiawithfourteethonoutermargin,thebasalone weaklydeveloped;foretibialspurexpandedandtruncateapically; foretibiae and forefemoralong and slender; forefemursmooth, finelypunctatethroughout.Metatibiaeobliquelytruncateapically; apicalspurspiniform.Abdomen.(Fig.2).Sternites3–7crenulately punctatealong theiranterior borders. Last abdominal segment slender.Pygidiumslightlyconvexandlightlyshagreenwith indis-tinctpunctuation;basalsulcusfineanddeepthroughout;marginal beadcontinuous.Malegenitalia.Parameres simple,taperingto apexinlateralview,apicalportionroundedin dorsalview.The internalsacoftheaedeaguswiththreehooks,twospine-likeand onespoon-like(Fig.4).

Allotype:Female.Totallength9.5mm.Maximumwidth5.0mm.

Sameasmalewiththefollowingsexualdifferences:clypealmargin anteriorly,notsobroadlyV-shaped;lateralpronotalmarginnot arched;lastabdominalsegmentbroadermedially;foretibiaeand forefemorashorterandforetibialspursslenderandslightlybentat tip;pygidiumnotaslong.

Variation:Totallength9.5–10mm.Maximumwidth5.0–5.5mm.

Materialexamined(34specimens):Holotype.Male:México:

Ver-acruz:DosAmates(Catemaco),8–V–1968,P.Reyes,M.Cabreracols. Nidodetuza.Cámaradedesechos(CEMT).Allotype.Female: ibi-dem,onefemale.Paratypes.ibidem,sixmales,fourfemales(three malesandthreefemalesatCEMT;onemale,GonzaloHalffter per-sonalcollection,Coatepec,Mexico;onemaleandonefemaleatBert Kohlmann personalcollection,Las Mercedesde Guácimo,Costa Rica;onemaletobedepositedatCanadianMuseum ofNature, Ottawa,Canada);onemale,onefemale,EntomologicalCollection attheInstituteofEcology,Xalapa,Mexico(IEXA), México: Oax-aca,Sta.MaríaChimalapas,12-VI-2015,colectadirecta,dentrode madrigueradetuza,J.LuisS.Huertacol.;fivemales,sevenfemales, ibidem,16-VI-2015;twomales,twofemales,ibidem,18-VI-2015; onemale,onefemale,ibidem,19-VI-2015;twofemales,ibidem, 22-VI-2015.

Remarks: This is the first North American inquiline Ateuchus

speciescollectedfromapocketgopherburrow.

Etymology:Thenametuza,anameinapposition,isthehispanized

commonname ofthe indigenousnahuatl wordtozan, givento thebeaver-lookingsubterraneanrodent,wherethenewAteuchus specieswasfound.

Geographicaldistribution:Thenewspeciesissofaronlyknown

fromthelocalityofDosAmates,Catemaco,inthestateofVeracruz, andthelocalityofSantaMaríaChimalapas,Oaxaca,Mexico.

Habitat:Thenewspecieswascollectedinthewastechamberof

apocketgopher(Rodentia:Geomydae)burrowduringthemonth ofMay.Althoughtherodentoftheoriginalcollectionsitewasnot

Figures4–5.A.tuzasp.nov.;4,drawingoftheinternalsac,lineequals1mm;5,A.

hornai(Balthasar,1939),dorsalhabitusofthefemaleholotype.

identifiedatthetime,itis mostlikely thatthespecimens were foundinsidethenestofOrthogeomyshispidus(LeConte).

Chorologicalaffinities:Interestingly,thisnewAteuchusspeciesis

foundundersimilarecologicalconditions,alowlandtropicalarea, asAteuchuscujuchiGénier,theotherknowninquilinerodent bur-rowAteuchusspeciesfromBolivia(Génier,2015).A.cujuchiwas collectedinaCtenomysburrow,arodentbelongingtoadifferent family,Ctenomydae,astheonecollectedinMexico.

Taxonomicrelationships:Theshapeofthebodyofthisspecies

resembles theSouth American species apicatum(Harold, 1867) andisclearlynotrelatedtotheknownbodyshapeoftheNorth orCentral AmericanAteuchusspecies.Thenewspecieswillkey tocouplet23/24(apicatum) inBalthasar’s(1939)key. Itcan be easilyseparatedfromA.apicatumbecauseofthepresenceofthe smallrugoseareaatthecenter ofthebaseoftheheadand the presenceofcoarsepuncturesatthemiddleofthepronotalbase. ThistaxonomicrelationshipwouldsuggestthatAteuchusspecies derivedofSouthAmericanlineshaveadaptedtoandcolonized low-landgophernestsinNorthAmerica;whereas,inthemountainous areas of North America it is species of the genus Onthopha-gus, which has adapted to colonizing gopher nests (Anduaga andHalffter,1991;LoboandHalffter,1994;ZuninoandHalffter, 2007).

Thisnewspeciesshowsadaptationstolivinginanenvironment devoidoflight,likethesubterraneangophernest.First,thedorsal eyeareaisverysmall.Second,theaforementionedrugoseareaat thebaseoftheheadisastridulationmechanism,similartothose foundatthebaseoftheheadofUroxys,asfoundinU.microcularis HowdenandYoung,U.microsBatesandU.platypygaHowdenand Young(DelgadoandKohlmann,2007;SolísandKohlmann,2013),

134 B.Kohlmann,F.Z.Vaz-de-Mello/RevistaBrasileiradeEntomologia62(2018)131–134

and mostprobablyhelps inthecommunication process of this species.

NewdistributionrecordsofandnotesonAteuchusWeberfrom

MexicoandCentralAmerica

Ateuchusguatemalensis(Bates),1887

ThisspecieshasbeenrecordedfromChiapas,Guatemalaand Hon-duras.ItishererecordedforthefirsttimefromNicaragua. Nicaragua: Jinotega: Cerro Kilambe, Camp 5, Las Torres, UTM–16P–1500283–0639500, 1200m, 23/30-IV-2001, col. J. SunyeryB.Hernández(5specimensatCEMT);Masaya:LasFlores, VII–1998,col.J.Téllez(1specimenatCEMT).

Ateuchus hornai (Balthasar, 1939), new combination (valid

species) (Fig.5)

ThisspecieswaspreviouslyconsideredasynonymofA.illaesum basedontheoriginaldescription,sincetheholotypewasnot avail-ableforstudyatthetime(Kohlmann,1984).Theexaminationofthe femaleholotypefoundintheNaturalHistoryMuseuminPrague, CzechRepublic(BezdekandHajek,2011),allowedustoconsiderita validspecies,distinguishablebythecharactersinthekeypresented herein.Wesuspectthetypelocality(Necaxa,Puebla,México)tobe wrong,andwehopethatamalewillbecollectedinordertoprepare amorethoroughdescriptionofthisinterestingspecies.

Conflictsofinterest

Theauthorsdeclarenoconflictsofinterest.

Acknowledgements

WethankDr.VinicioSosafromtheInstituteofEcology,Xalapa, Mexico,forhisinformationregardingthepossibleidentityofthe rodent,wherethenewAteuchusspecieswascollected. Wealso thanktheAlexander vonHumboldt Foundation,who most gra-ciouslydonatedtheopticalequipmentusedinthisstudy,aspart ofthesupportgiventothefirstauthor(B.K.),duringhis sabbati-calstayattheTechnicalUniversityofDresdenin2010.Wethank JiriHajek,NationalMuseum,Prague,forthekindnessof prepar-ingthe pictures of theholotype ofChoeridium hornaithat was examinedbyF.Z.V.M.duringa sabbaticalyear2013–14 granted by CNPq (202327/2013-2), with one trip to Prague funded by F.Z.V.M.isaCNPq(ConselhoNacionaldeDesenvolvimentoCientífico

eTecnológico)fellow(302997/2013-0and306745/2016-0)andis grantedbyFAPEMAT(Fundac¸ãodeAmparoàPesquisadoEstadode MatoGrosso(PRONEM568005/2014)andCNPq(405697/2013-9, 440604/2015-0)).B.K.hadthepossibilitytovisitCuiabáinBrazil thankstoaFAPEMAT(Fundac¸ãodeAmparoàPesquisadoEstadode MatoGrosso)grantforimprovementoftheinternationalizationof thePostgraduatePrograminEcologyandConservationof Biodiver-sityofMatoGrossoFederalUniversity(UFMT).Lastbutnotleast, wethankKentMcLeodforrevisingtheEnglishlanguage.

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